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Forest Ecology and Management

journal homepage: www.elsevier.com/locate/foreco

Deadwood management in Central European forests: Key considerations forpractical implementation

Lucie Vítková⁎, Radek Bače, Petr Kjučukov, Miroslav SvobodaDepartment of Forest Ecology, Faculty of Forestry and Wood Sciences, Czech University of Life Sciences, Kamýcká 129, Praha 6 – Suchdol 16521, Czech Republic

A R T I C L E I N F O

Keywords:Long-term retentionSaproxylic diversityMicrohabitatsForest management

A B S T R A C T

A substantial amount of literature on the importance of deadwood in Central European forests has been availableproviding partial recommendations to enhance deadwood-dependent biodiversity. However, a comprehensivereview of science- and forestry experts-based recommendations effectively enhancing deadwood bearing in mindoperational implications has not been presented in international literature. Therefore, this paper compiles thekey aspects regarding the implementation of deadwood management in managed forests where the aim is tofavour biodiversity without compromising or negatively affecting operational and commercial aspects of forestmanagement. Simple deadwood management guidelines rooted in science and forestry expertise aiding decision-making in the efforts to effectively enhance biodiversity without compromising other management objectives arethus provided. Specifically, long-term retention of individual trees or tree groups and the retention of alreadyexisting deadwood (e.g. snags, coarse woody debris, uprooted, snapped, and sun-exposed trees) as well as ar-tificial creation of deadwood (e.g. tree girdling) are presented here as we identified them as the key approachesto successful deadwood management. The major advantages and disadvantages of individual deadwood man-agement approaches in terms of biological and operational/commercial aspects are also emphasised in order toassist forest managers in their decision-making. Furthermore, the key factors that should be considered whenapplying ecologically and economically efficient deadwood management are discussed; i.e. retention of treeswith microhabitats, size of retained trees, position and arrangement, and decay stage. The main points regardingthese factors are also addressed in the light of supporting realistic implementation of individual deadwoodmanagement approaches.

1. Introduction

Biodiversity is considered a fundamental driver of high intrinsicvalue that steers forest ecosystem functionality and facilitates for keyecosystem processes and services (Mori et al., 2017). An increasingamount of evidence supporting the significance of deadwood for bio-diversity has been available. Although the significance of deadwood asa support for biodiversity has been widely recognised (e.g.Vandekerkhove et al., 2005; Bütler et al., 2007; Lassauce et al., 2011;Lachat et al., 2013; Bouget et al., 2014a, 2014b, etc.), deadwood wasalso reported to be important for carbon storage (Kueppers et al., 2004;Woodall and Liknes, 2008; Olajuyigbe et al., 2011), nutrient cycling(Laiho and Prescott, 2004; Yuan et al., 2017), soil forming processesand hydrology (Harmon et al., 1986), etc. Deadwood volumes in forestsgreatly vary depending on forest type (Christensen et al., 2005), treespecies (Debeljak, 2006), stand age (Ekbom et al., 2006), geographical

location (Stokland et al., 2012) as well as other factors. However, forestmanagement and natural disturbance history also alter the volume ofdeadwood as well as its type and distribution throughout the forest.

Deadwood is generally present in rather low volumes in con-ventionally managed forests in comparison to natural forests (Siitonenet al., 2000; Pedlar et al., 2002; Debeljak, 2006; Larrieu et al., 2012;Dieler et al., 2017; Nagel et al., 2017). This is mainly due to the har-vesting of trees once they reach the target diameter for felling, whichallows to retain only a small amount of deadwood typically in a form ofshort stumps, small twigs and branches resulting in the absence of snagsor large logs (Kruys et al., 1999). However, larger segments of dead-wood are particularly important as they remain longer in the forestecosystem continuously providing habitat as opposed to deadwood ofsmall dimensions offering habitat only temporarily (Lachat et al.,2013). It is also essential to manage for diversity in the retaineddeadwood; i.e. a range of sizes, decay stages, tree species, locations, etc.

https://doi.org/10.1016/j.foreco.2018.07.034Received 19 April 2018; Received in revised form 17 July 2018; Accepted 18 July 2018

⁎ Corresponding author at: Department of Forest Ecology (Room 409), Faculty of Forestry and Wood Sciences, Czech University of Life Sciences, Kamýcká 129,Praha 6 – Suchdol, 165 21, Czech Republic.

E-mail address: [email protected] (L. Vítková).

Forest Ecology and Management 429 (2018) 394–405

0378-1127/ © 2018 Elsevier B.V. All rights reserved.

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in order to provide suitable environment for a variety of deadwood-dependent species. Deadwood in natural forests results from tree mor-tality caused by senescence processes or by competition. Alternatively,deadwood may be created by natural disturbances whose quantity andtype is rather variable (Rahman et al., 2008) similarly as the deadwoodtypes and quantities it creates (e.g. splintered stems, snapped or brokenstems and branches, uprooted trees, etc.). The structure and function ofthe deadwood changes over time following the natural disturbances(McComb and Lindenmayer, 1999) also depending if any silviculturalinterventions are consequently applied. Salvage logging, for instance,tends to remove most of the deadwood substantially reducing theoverall deadwood quantities (Priewasser et al., 2013, Michalová et al.,2017).

To a limited extend, the concept of deadwood management has beena part of forest management of some state forest enterprises as well asfor some private forest properties in e.g. Germany, Switzerland, France,Italy, Austria, Sweden and Denmark. Life and Life+ projects reportsalso provided relevant information on some of the practical cases ofdeadwood management (i.e. Cavalli and Mason, 2003; Mason et al.,2003) with NATURA 2000 reports also delivering information ondeadwood management in some of the above-mentioned Europeancountries (European Commission, 2015). Nonetheless, outcomes offorest management practices focusing on the enhancement of dead-wood quantity and quality have not been available in published inter-national literature with only some exceptions (e.g. Doerfler et al. (2017)focusing on the success of a deadwood enrichment strategy in pro-duction forests in Germany). Although deadwood volumes are assessedon a 5-years basis across European Union (e.g. Forests Europe, 2015),detailed information on deadwood quantities of different types andstages of decay in production forests is unavailable in many countriessince the management practices aiming at deadwood enhancementhave been applied only for the last 20 years. This period is too short toallow us to record sufficient data on the development of deadwood overtime. Besides, detailed deadwood monitoring that would normally yieldvaluable data on deadwood quantities and qualities has not been part ofnational forest inventories in some countries. However, although stra-tegies to increase the deadwood qualities in managed forests have beenimplemented in the light of biodiversity enhancement and certifiedsustainable forest management (e.g. PEFC, 2010; FSC, 2013) and theyhave been monitored as a part of National Forest Inventories in somecountries (e.g. Germany, Switzerland, the Netherlands), their successhave been largely under-reported across Europe (Doerfler et al., 2017).Nonetheless, deadwood has been stated as one of the indicators ofsustainable forest management.

Maintenance of sufficient deadwood quantities comprising of avariety of deadwood types does not only locally increase saproxylicspecies diversity. It also reduces the risk of the saproxylic species be-coming extinct thanks to the presence of suitable habitat and thus theirviable population. Maintaining sufficient population also reduces therisk of undesirable loss of genes, which occurs during prolonged re-duction in population due to ecological and stochastic reasons (Øklandet al., 1996). A variety of deadwood types shall be also encouraged,with the same applying to the diversity of deadwood’s spatial dis-tribution and stages of decay supporting a range of habitats since dif-ferent species require different conditions (Bouget et al., 2013). Theimportance of the presence of old trees and continuous deadwoodsupply in the conservation of red-listed species was also highlightedespecially since the occurrence of relict saproxylic species correlateswith the continuity in forest cover containing such features (Buse,2012). The retention of trees bearing microhabitats and deadwoodshould be thoughtfully planned in order to ensure long-lasting habitatcontinuity (Bütler et al., 2013).

A substantial amount of literature on various aspects of deadwoodproviding and summarising valuable information on deadwood di-versity and volume but also the importance of deadwood for ecology ofsaproxylic species has been available among other topics (Heilmann-

Clausen and Christensen, 2003; Heilmann-Clausen et al., 2005; Müllerand Bütler, 2010; Lassace et al., 2011; Bouget et al., 2012, 2013;Dittrich et al., 2014; Müller et al., 2015a; Gossner et al., 2016; Doerfleret al., 2017). Individual recommendations supporting specific dead-wood management approaches have also been offered; i.e. the necessityto have certain deadwood quantity and/or quality (dimension, position,tree species, etc.) (e.g. Kappes et al., 2009; Müller and Bütler, 2010;Doerfler et al., 2017). However, international literature conciselycompiling and presenting a combination of science- and expertise-basedrecommendations guiding the enhancement of deadwood volumes anddiversity in managed temperate forests of Central Europe has not beenavailable; especially when considering commercial feasibility of in-dividual approaches but still bearing in mind the biodiversity en-hancement. Although deadwood-related literature often concludes withrecommending the increase in deadwood volumes and/or deadwooddiversity (e.g. Bunnell and Houde, 2010; Müller and Bütler, 2010), arange of specific, simple and feasible operational approaches that canbe implemented to achieve these deadwood management re-commendations is rarely mentioned. Especially, if the balance betweenthe deadwood management benefits biodiversity with operational andcommercial aspects also taken into an account.

Therefore, the major aim of this paper is to concisely present the keydeadwood management approaches – based on scientific findings andexpertise - that can be considered in public and private temperateforests of Central Europe. This is conducted in the efforts to effectivelyenhance deadwood volume and its diversity without compromisingother management objectives or increasing operational costs. The majorfactors necessary to be considered when applying an effective dead-wood management are also included in order to support its realisticimplementation in practice. We further emphasise the major benefitsand drawbacks of individual deadwood management approaches inorder to provide a representative picture of its application bearing inmind operational feasibility and commercial viability of these ap-proaches.

2. Approaches to forest management enhancing deadwood

Based on the vast amount of literature published on deadwood as animportant biodiversity indicator as well as on existing expertise, severalapproaches that can be used to increase deadwood quantities and typesin managed forest were identified; i.e. the retention of single trees orgroups of live trees or the retention of snags and already existingdeadwood. Although these methods reflect research concerning thefunctional effectiveness of measures to promote biodiversity (e.g.Fedrowitz et al., 2014; Hämäläinen et al., 2014), other methods such asretention of lying logs following harvesting, retention of uprooted treesor artificial creation of deadwood by generating high tree stumps orkilling of targeted trees (e.g. girdling) can also be considered. Combi-nation of individual deadwood management approaches is also con-sidered a suitable concept for deadwood management in order toachieve greater volume and diversity of deadwood (Ranius et al.,2005).

Opting for deadwood enhancement approaches can increase thediversity (decay stage and dimensions) of deadwood, which is moreimportant for biodiversity than the actual deadwood quantity (Rimleet al., 2017). It is important to work with natural processes that createdeadwood but also to improve linkages between existing deadwoodfeatures by artificially generating additional deadwood as well as pro-tecting already existing deadwood (Humphrey and Bailey, 2012).Deadwood management is challenging and presents numerous trade-offs between biodiversity enhancement and operational or commercialaspects. Therefore, some of the major advantages and disadvantages ofselected deadwood management approaches forest managers are likelyto encounter when adopting deadwood management are demonstratedin Tables 1 and 2. Although some efforts towards tree retention alreadytakes place in many commercial forests, it is important to bear in mind

L. Vítková et al. Forest Ecology and Management 429 (2018) 394–405

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that the resulting deadwood volumes should be within a recommendedrange considering the state of forest stand, forest management history,species composition, stand age, etc. (e.g. Müller and Bütler, 2010). Inthe following sections, we present some of the major practices that canbe employed to enhance deadwood that, at the same time, reflect de-cisions likely faced when adopting effective deadwood management.

2.1. Retention of living single trees and tree groups

Allowing individual trees or small groups of trees to reach the end oftheir life span is an effective measure to increase the amount of dead-wood. The retained individual trees or tree groups can reach the stageof veteran trees that eventually develop into snags (unless damaged bywind and consequently fallen on the ground) that decay disintegrating

Table 1The main advantages and disadvantages of major biological aspects of selected deadwood management approaches aiming at deadwood enhancement in managedforests.

Management approach Advantages Disadvantages

Individual (live) treeretention

Creates important stepping stones and microhabitat for saproxylicbeetles; provides key microhabitat for future development of standingand consequently lying deadwood; provides continuous supply of seeds;supports the development of old-growth elements in the forests

If retained trees are too small, once decayed, they may not form asuitable habitat for saproxylic species requiring larger segments ofdeadwood

(Live) tree group retention Creates important microhabitat for saproxylic fungi or epixylicbryophytes; forms buffering effect - microclimate; provides keysubstrate for future development of standing and consequently lyingdeadwood; provides continuous supply of seeds; supports thedevelopment of old-growth elements in the forests

If retained trees are too small, once decayed, they may not form asuitable habitat for saproxylic species requiring larger segments ofdeadwood

Retention of existingdeadwood

Already existing deadwood of advanced stages of decay providesmicrohabitats for saproxylic organisms; supports the development ofold-growth elements in the forests

May be a source of pests in the case of susceptible species (e.g. Norwayspruce and bark beetle); freshly created snags may require longer timeto become suitably decayed

Retention of uprooted trees Provides microhabitats for saproxylic species; supports the developmentof old-growth elements in the forests

Freshly uprooted trees require longer time to become suitably decayedfor some saproxylic species; possible entry point or a source of pests incase of susceptible species (e.g. Norway spruce and bark beetle)

High stump retention/creation

Provides suitable substrate for the development of short snags andmicrohabitats for saproxylic organisms

Depending on the height of the stump, the decay and disintegration canbe rapid

Live trees girdling Supports the creation of deadwood and consequently microhabitats forsaproxylic organisms; supports the development of old-growth elementsin the forests

Girdled area may be an entry point for unwanted tree pests and diseases

Retention of deadwood ofdesired tree species

Supports (not only) specialist saproxylic species bound to deadwood ofparticular tree species

Potential source of pests in the case susceptible species are retained (e.g.Norway spruce and bark beetle)

Retention of trees of specificdiameter

Creates substrate for the development of deadwood of dimensionssuitable for (not only) specialist saproxylic species bound to deadwoodof specific tree species; supports the development of old-growthelements in the forests

If the retained trees are too small, saproxylic species requiring largedeadwood segments may not be able to colonise (and vice versa);retention of small trees results in their rapid decay

Retaining sun-exposed treesand deadwood

Creates substrate for the development of deadwood with suitableproperties for specialist saproxylic species requiring warm and sun-exposed conditions; supports the development of old-growth elementsin the forests

If the retained trees are too small, saproxylic species requiring largedeadwood segments may not be able to colonise (and vice versa);retention of small trees results in their rapid decay

Table 2The major operational/commercial aspects of individual management approaches aiming at deadwood enhancement in managed forests where the main advantagesand disadvantages are presented.

Management approach Advantages Disadvantages

Individual (live) tree or treegroup retention

Reduces harvesting and logging cost if the retained trees are located inareas with difficult access (extreme slopes, rugged terrain, etc.)

Potential loss of financial revenue as the retention of individual treesor tree groups may involve trees of larger diameters aimed at timberproduction

Retention of existingdeadwood

Reduces harvesting or tending costs as decaying trees are of low or nocommercial value for timber or fuel wood production or are located in anarea with difficult access

May form obstacles to logging operations or to access routes or onroads and paths, etc.; possible health and safety risk to public if closeto roads, paths, etc.

Retention of uprooted trees Reduces harvesting or tending costs as uprooted trees are of lowcommercial value for timber production or located in an area with difficultaccess

May pose health and safety risk to forest workers or may create anobstacle when located close to roads, paths, etc.

High stump retention/creation

Creating high stump from a tree of poor form at its base does not decreasequality of the harvested log; easier in the mountain forests

Additional costs involved as trained and skilled staff is necessary tocarry out the operation

Live trees girdling If poor quality trees are retained for girdling, the overall quality ofharvested timber is not decreased. Could be used to kill non-native species

Additional costs involved as trained and skilled staff is necessary tocarry out the operation especially if carried out repeatedly acrosslarger areas

Retention of deadwood ofdesired tree species

Undesirable species for timber production are retained to reduce theharvesting and transport costs especially if located in an area with difficultaccess

Possible loss of financial revenue as the retention may involve treesaimed at timber production

Retention of trees of specificdiameter

Trees of undesirable dimensions are retained, which reduces theharvesting and transport costs, especially if located in an area withdifficult access

Potential loss of financial revenue since the retention of individualtrees of larger dimensions may be aimed at timber or fuel woodproduction

Retaining sun-exposed treesand deadwood

Retention of poor quality trees or those located in an area with difficultaccess that are exposed to sun reduces the harvesting and transport costs,especially if located in an area with difficult access

If under denser canopy, heavier thinning may be required in theclose proximity of the retained tree(s) to create more light accessingthe tree


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into coarse woody debris with progressing time. The decay rate in-creases with e.g. rising air humidity and temperatures or the length ofvegetative period. Priority should be given to the oldest and largesttrees, preferably those with already existing microhabitats (for detailsee Section 3) and with some extent of deadwood (Bouget et al., 2014a,2014b; Müller et al., 2014). When choosing trees to be retained and leftfor snag formation, trees with the presence of microhabitats or defectsfound within the first 2 m of the bole should be prioritised. Healthy logsand stems with the diameter< 30 cm should be used for retention onlyif there are no other options since it is beneficial to keep larger seg-ments of deadwood due to their longer residence time (Bače andSvoboda, 2015).

There are several options in tree retention; the group of trees orindividual trees can be marked and retained in production forests withthe remaining stand being subjected to usual silvicultural interventions.Alternatively, trees can be located at more open site with direct lightreaching the retained trees. An area between the forest stand and ameadow, field, water body, etc. may be also considered a suitable placewithout the necessity to excessively open the canopy and thereforeminimise the loss of timber production and quality of future standswithin the area. It is possible to use deforested areas in continuouslywooded landscapes, such as electricity pylon routes, abandoned foresttracks, rides or pastures, etc.

A combination of dispersed and aggregated patterns can be usedwhen retaining individual trees and tree groups (Bütler et al., 2013).The retention of aggregated tree groups was reported to provide sui-table sites for birds than in the case of dispersed retention (Bütler et al.,2013). Similarly, lichens perform better within a small group of re-tained trees as opposed to dispersed individuals (Nascimbene et al.,2013). Thorn et al. (2017) reported that this is the case for the majorityof saproxylic species. The retention of sun-exposed scattered trees, onthe other hand supports umbrella species such as great Capricorn beetle(Cerambyx cerdo L.) (Buse et al., 2007; Albert et al., 2012) or long-horned beetle (Rosalia alpina L.) (Russo et al., 2011).

2.2. Retention of existing deadwood

If deadwood of various decay stages and sizes already exists at a siteof interest (e.g. snags, fallen deadwood or uprooted trees), it is fa-vourable to preserve it. If any standing dead trees or snags of preferablylarge dimensions are available, they should be retained in full.However, exceptions may include snags that pose an imminent risk tothe health and safety of public (i.e. snags near publicly accessible roadsand paths, places with increased recreational activities) and forestworkers (tending operations, marking, etc.) or if they form a majorobstacle for harvesting. Nonetheless, if a snag needs to be felled, itsremnants should be left where the felling took place, if possible.

Similarly, as in the case of snags, as far as possible, already existinglying deadwood shall be retained. Lying deadwood resulting fromnatural disturbances (uprooted trees and trees broken by wind/snow)tend to be subjected to salvage logging. Should deadwood resultingfrom natural disturbances pose no risk to the production forests in termsof generating greater pest outbreaks, some of the uprooted or snappedtrees can be retained. Uprooted trees create favourable conditions fornatural regeneration in the form of exposed mineral soil and pit-and-mound topography, while being important for natural soil development(Šamonil et al., 2010, 2014). Displaced root plates should be preventedfrom returning to their initial position following the disturbance. Log-ging practices (careless winging or skidding) damaging already existingdeadwood or uprooted trees should be also prevented.

2.3. Artificial creation of deadwood

Retaining thick crown branches or entire crowns after felling op-erations is considered a suitable tool to increase the amount of lyingdeadwood along with the retention of poorer quality stems after

harvesting (Doerfler et al., 2017). Implementation of such deadwoodenhancing strategy is considered an achievable way to promote biodi-versity in the production forests (Doerfler et al., 2017). Uprooted trees,high stumps, snags as well as standing and leaning dead trees can becreated artificially (e.g. Cavalli and Mason, 2003; Nordén et al., 2004;Brin et al., 2013; Mason and Zapponi, 2015). Active creation of thesedeadwood features usually include tree girdling, felling and pulling,inoculation with fungal pathogens or combination of these techniquesby skilled chainsaw operators (Lewis, 1998; Kuuluvainen et al., 2004).

Trees with microhabitats also bearing deadwood can be generatedusing some of the above-mentioned approaches along with de-limbingand active creation of cavities and basal pockets of various sizes (Lewis,1998). The nesting cavities can be created by cutting and extracting awood block from the tree 1–4m above the ground and cutting a thinslice to make a lid to close the cavity (Zapponi et al., 2015). Basalpockets, on the other hand, can be made by creating slits using achainsaw (Zapponi et al., 2015). Pollarding (i.e. the periodical removalof the upper branches of a tree by pruning; Thomas, 2000) was reportedto lead to the rapid formation of cavities and is thus considered animportant technique in cavities-dependent species and saproxilic ha-bitats restoration (Sebek et al., 2013). Other microhabitats such as stemsplintering and various injuries further developing into microhabitatscan be also generated using a chainsaw.

Beside additional costs and the necessity to employ specially trainedstaff the above-mentioned practices may be limited by legislation insome countries. However, the benefits of these deadwood-enhancingpractices also leading to creation of crucial microhabitats and thusgreatly facilitating for biodiversity are likely to offset the costs andefforts involved.

3. Key factors to be considered as a part of deadwoodmanagement

A range of factors affects deadwood management causing differ-ences in the trends of species diversity in different species groups. Inorder to support the key factors that shall be considered in deadwoodmanagement, selected key factors are presented in Table 3 in order todemonstrate the differences in the trends of selected deadwood-de-pendent species groups such as lichens, bryophytes, fungi, beetles,vertebrates and invertebrates.

3.1. Retaining trees with microhabitats

If available, trees harbouring microhabitats shall be favoured forretention when making a decision on retaining a group of trees or in-dividual trees (Bouget et al., 2014a, 2014b; Müller et al., 2014; Fig. 1,Table 3). Trees with microhabitats often bear some amount of dead-wood providing necessary habitats for a range of species (or groups ofspecies) to grow, nest or forage as a part of their life cycle (Winter andMöller, 2008; Vuidot et al., 2011; Larrieu et al., 2014; Paillet et al.,2017; Larrieu et al., 2018; Kozák et al., 2018). The retention focus shallbe also made on individuals with the potential to develop fully func-tioning microhabitats over time (e.g. presence of injuries, harvestingdamage, etc.). Such trees shall be selected early; if necessary, theyshould be released from competition by means of thinning to preventsuppressed crown and root system ensuring the full development ofmicrohabitats and deadwood (Krása, 2015). If information is availableon individual’s age, attempts should be made to give the priority to theoldest trees as they often feature well-developed microhabitats (Winterand Möller, 2008; Michel and Winter, 2009; Vuidot et al., 2011).

The number of trees bearing microhabitats is minimised in pro-duction forests since conventional silvicultural approaches remove treeswith microhabitats as such trees are considered of poor quality in termsof timber production. The trees bearing microhabitats are rarely fullydeveloped in production forests due to relatively short rotation periodsand harvesting of trees once they reach target diameter as opposed to


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leaving them to reach the end of their life span as in natural forests. Thiswas confirmed by studies comparing the abundance of microhabitats inproduction and natural unmanaged forests (Winter and Möller, 2008;Vuidot et al., 2011; Regnery et al., 2013; Paillet et al., 2017). Althoughmicrohabitats are found in production forests, their diversity is usually

rather limited to e.g. bark loss, dendrothelms, cavities, etc. (Larrieuet al., 2012). However, it is still favourable to focus on retention of treesthat bear such microhabitats. In addition, the retention of trees of dif-ferent sizes, species and decays stages that bear microhabitats shall beconducted at a range of sites; i.e. sites with special designation/

Table 3Trends of the differences in species diversity given different deadwood factors; figures represent the proportion of particular taxon in relation to the maximumnumber of species given the level of particular factor - the scales of individual parameters were united to simplify their interpretation. The description of decay stagesis detailed in Appendix A.

Taxon Factors

Stage of decay Position Mean size Light exposure Tree species

0 1 2 3 Standing Lying 10–20 cm >20 cm Direct light Semi-shade Full shade Conif. Broadl.

Lichens1,2 0.40 0.73 0.88 0.38 1.00 0.25 0.55 1.00 0.80 1.00 0.80 – –Bryophytes1,2 0.13 0.63 0.70 0.65 0.00 1.00 0.65 1.00 0.10 0.50 1.00 –Fungi1,2 0.33 0.70 0.80 0.73 0.30 1.00 0.80 1.00 0.25 0.80 1.00 –Beetles2,3,4 0.70 1.00 0.85 0.10 1.00 0.70 0.95 1.00 0.97 0.95 0.47 –Vertebrates1 1.00 1.00 0.75 0.30 – – – – – – – – –Invertebrates4 0.30 0.40 0.70 0.95 – – 0.85 1.00 1.00 0.80 0.80 – –All species5,6 0.45 0.88 0.90 0.35 0.83 0.95 0.75 1.00 1.00 0.15 0.35 0.60 1.00Weighted

average0.50 0.88 1.00 0.67 0.80 1.00 0.80 1.00 1.00 0.60 0.69 0.60 1.00

Trend (weightedaverage)

1 Bunnell and Houde (2010).2 Jonsson et al. (2010).3 Lindhe et al. (2005).4 Jonsell et al. (2004).5 Stokland et al. (2004).6 Stokland and Meyke (2008).

Fig. 1. Key factors of deadwood management along with the major points worth bearing in mind when considering deadwood management.


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protection status, commercially productive as well as unproductivesites, different forest types, diverse altitudes, etc.

3.2. Size of retained deadwood

Enhancing the occurrence of deadwood of large dimensions wasreported to have positive effects on biodiversity (e.g. Økland et al.,1996; Gossner et al., 2013; Svensson et al., 2014; Juutilainen et al.,2014; Seibold et al., 2014). The presence of large segments of dead-wood is considered a more important factor in comparison to, for ex-ample, the position of deadwood (standing/lying) in explaining theoccurring species diversity (Bouget et al., 2012) (Fig. 1, Table 3). Largesegments of deadwood are also favoured since greater tree diametercorrelates with thicker bark with diverse features (e.g. increased levelof cracks and rugged texture of the outer bark) (Bače and Svoboda,2015) or a long-term provision of nutrients (Herrmann and Bauhus,2018). Larger dimensions contribute to longer decay time and thuslonger period for the suitable habitat to be available (Heilmann-Clausenand Christensen, 2004). In addition, deadwood of greater volume has asmaller surface area/volume ratio, which results in a greater stability oftemperature and moisture in deadwood (Bače and Svoboda, 2015).

Large logs cannot be replaced with the equal volume of smaller logsin the light of supporting the enhancement of biodiversity since manyspecies only require deadwood of larger proportions (Kraus andKrumm, 2013). Although the retention of high stumps was reported tobe more beneficial in comparison to brash retention, the combination ofboth brash and high tree stumps retention was stated as a suitableoption in terms of economy and functional effectiveness for biodiversity(Ranius et al., 2014). The presence of deadwood of large dimensionscan be considered as a good indicator of continuity since their wooddecomposition is rather slow. The continuity of deadwood is of highimportance as it facilitates for better connectivity and allows organismsto better disperse, interact, and access resources (North and Keeton,2008).

Unlike deadwood of large dimensions, deadwood of small dimen-sions in a form of small branches and small stems can be found inproduction forests (Fridman and Walheim, 2000; Bače and Svoboda,2012). If forest management measures incorporating deadwood en-hancement are adopted, it is necessary to enhance the diversity ofdeadwood dimensions and actively focus on the development of largesegments of deadwood. However, smaller deadwood segments are alsonecessary since some saproxylic species require smaller stem or branchthickness (Ódor et al., 2006; Juutilainen et al., 2014).

3.3. Tree species

It is appropriate to retain native species whose occurrence is sparse(Økland et al., 1996) in order to support local species communities andthus achieve a greater diversity in deadwood of native species compo-sition. Tree species with slow rate of decay should be prioritised forretention since the rate of decay varies amongst individual species; oaks(Quercus spp.) have a slower decomposition rate in comparison tospruces (Picea spp.), pines (Pinus spp.), and European beech (Fagussylvatica L.) that decompose 1.4, 1.6 and 1.8 times faster than oaks,respectively (Rock et al., 2008). Moreover, broadleaved species tend tobear more microhabitats, which should be considered when choosingwhich trees to retain (Larrieu et al., 2012). It is important to includelate successional species such as European beech as well as pioneerspecies e.g. birch (Betula spp.), poplar (Populus spp.), and willow (Salixspp.) where possible (Fig. 1, Table 3).

Although it is also important to enhance deadwood in coniferousforests, conifers tend to support less microhabitats, which is the case forvarious geographical locations in Europe. This is due to multiple rea-sons such as natural lack of certain microhabitats in conifers due toevolution in the mountain areas, their use in plantation forestry wherethey rarely reach the senescence age when microhabitats are formed,

etc. More microhabitats were reported on European beech in compar-ison to silver fir (Abies alba Mill.) in beech-fir forests in central Pyreneeswhere fir bears no microhabitats below the diameter of 68 cm (Larrieuet al., 2012). In Scandinavian forests, deadwood of broadleaved speciesgenerally supports a greater diversity of saproxylic species than dead-wood of coniferous species (e.g. Stokland et al., 2004). The oak genusshould be emphasized in particular; pedunculate (Quercus robur L.) andsessile oaks (Quercus petraea (Matt.) Liebl.) are important for biodi-versity of saproxylic invertebrates in Central Europe (Vodka et al.,2009; Bouget et al., 2011). In Sweden, for instance, 32% (i.e. 174) ofRed-listed saproxylic species are bound to oak species (Jonsell et al.,1998). In the case of Central Europe, many saproxylic beetles are at-tracted to hornbeam (Carpinus betulus L.) in the first years of its wooddecomposition, even if left in the shade (Müller et al., 2015b). In con-trast to hornbeam, although also supporting some specialised lichenand moss species, ash (Fraxinus excelsior L.) trees do not host that manysaproxylic species, which is perhaps linked to specific chemistry of itswood (Müller et al., 2015b). Sycamore (Acer pseudoplatanus L.), how-ever, another broadleaved tree species, is significant for the biodiversityof lichens, particularly at higher elevations (Bače and Svoboda, 2015).

3.4. Position and arrangement

The position of deadwood influences its properties since many fungiand bryophytes are favoured by shaded moist conditions while dry andwarmer conditions are generally suitable for saproxylic beetles and li-chens (Lachat et al., 2013) (Fig. 1, Table 3). Therefore, it is important toretain sun-exposed trees in gaps or under more open canopy as a part ofdeadwood enhancing management (Rosenwald and Löhmus, 2008;Gustafsson et al., 2010; Parmain and Bouget, 2018) as such featuressupport valuable microhabitats favouring saproxylic umbrella species(Buse et al., 2007; Albert et al., 2012; Russo et al., 2011). Moreover, thesun-exposed and warm conditions in gaps may appeal to shade-intol-erant beetle species that are attracted by the openness as opposed to theactual microhabitat trees bearing deadwood (Koch Widerberg et al.,2012). Deadwood with elevated moisture, on the other hand, showedgreater fungal community succession, which was especially pronouncedin advanced stages of decay (Rajala et al., 2012).

It is necessary to consider diversity of deadwood types when re-taining particular trees or tree groups in order to correctly meet thedeadwood management objectives. In some cases, standing deadwoodhosts more saproxylic species than lying deadwood as demonstrated one.g. oak snags that facilitate a higher number of individuals of sa-proxylic beetles than lying logs (Bouget et al., 2012). Lying deadwoodon the other hand tends to host a greater number of fungi and bryo-phytes in comparison to standing deadwood (Jonsson et al., 2010). It isalso important to note that similar substrates at different locations, e.g.forest floor, tree stem or canopy branches, can facilitate conditions fordifferent species; snags, for instance, accommodated for species thatwere absent from lying logs (Bouget et al., 2012).

3.5. Decay stage

Saproxylic species richness and the occurrence of rare saproxylicspecies is determined by the presence of deadwood that is, amongstother factors such as deadwood size, determined by the decay stage(Heilmann-Clausen and Christensen, 2004; Sefidi and Etemad, 2015).Therefore, enhancing diversity of deadwood types in different stages ofdecay (retention of existing deadwood and creation of new segments) isconsidered a viable management option to increase biodiversity inforests (Ódor and Standovár, 2001) (Fig. 1, Table 3). Decaying woodhosts a large number of lichens, bryophytes, fungi and invertebrateswhose occurrence is influenced by the decay stage (Ódor andStandovár, 2001; Ódor and van Hees, 2004; Penttilã et al., 2004;Siitonen et al., 2000). Intermediate and advanced decay stages appearto be the most favourable for many species of fungi, with the more


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progressed intermediated stage being particularly favourable for someRed-listed species (Heilmann-Clausen and Christensen, 2003; Persianiet al., 2015; Sefidi and Etemad, 2015). Small vertebrates, on the otherhand, seek earlier stages of decay for foraging (Bunnell and Houde,2010).

Different deadwood types vary in their decay dynamics; i.e. thedecay rates of standing deadwood tend to be slower than in the case oflying deadwood (Jomura et al., 2008; Bouget et al., 2012). It is thusnecessary to retain or create different types of deadwood as part offorest management to support a range of decay stages that host avariety of not only saproxylic species. Apart from its significance forsaproxylic diversity, decaying wood also supports natural regenerationespecially in mountain forests as deadwood as a substrate favours theoccurrence and survival success of natural regeneration of e.g. Norwayspruce (Zielonka, 2006; Bujoczek et al., 2015). A study from ŠumavaNational Park (south-eastern Czech Republic) showed that mostNorway spruce seedlings grew on lying deadwood of intermediate andadvanced decay stages (Čížková et al., 2011). This was attributed todiverse and rugged surface of the deadwood but especially to the stablehydrological conditions and better accumulation of water in the dead-wood offering suitable microhabitat conditions for Norway spruceseedlings.

4. Suitable settings for an effective deadwood management

Individual forests comprise of diverse site conditions of differentvalue when initiating deadwood management. Therefore, several con-siderations need to be taken into an account when planning the ap-plication of deadwood management. Prior to the deadwood manage-ment application, it is important to consider several aspects in the lightof assessing the site’s suitability to particular deadwood enhancingapproach: (i) current levels of deadwood on site, (ii) continuity anddiversity of deadwood habitats over time, (iii) interest in conservationof specific saproxylic, (iv) ecological connectivity and (v) history ofmanagement (Humphrey and Bailey, 2012).

It is important to preserve deadwood in parts of forests of higherecological value where deadwood already occurs. Nonetheless, dead-wood should also be enhanced in areas where it is absent since even asmall increase in deadwood quantity may be a positive prerequisite forsome saproxylic species to occur. The implementation of different ap-proaches aiming at deadwood enhancement should ensure unevendistribution of deadwood across the forest with the efforts to increasethe proportion of deadwood in places where it is absent. However, thedecision-making should be case-specific due to a large variation in site

conditions; an example of the potential to implement deadwood man-agement is demonstrated in Fig. 2; a case study of a forest propertycommonly found in Central Europe.

Natural forests are, by definition, rich in deadwood, in comparisonto production forests, and may serve as refugia of specialist saproxylicspecies that have an increased demand on the amount of deadwood(Gossner et al., 2013). Forest reserves or forests stands where har-vesting was abandoned were also studied to show the growing amountsof deadwood providing valuable habitats (Vandekerkhove et al., 2009;Meyer and Schmidt, 2011; Paillet et al., 2015). If there are natural re-serves around the area where higher quantities of deadwood alreadyoccur, priority should be given to retaining deadwood near such sites inorder to create ‘stepping stones’ and thus support suitable conditions forthe populations of saproxylic species and their potential to spread fur-ther. In order to create similar deadwood hotspots in managed forests, itis favourable to focus the deadwood enhancing practices in areas notdesired or suitable for timber production such as waterlogged depres-sions, rocky slopes and buffer zones along watercourses. Refrainingfrom timber harvesting in such places or in difficult terrain also reducesthe harvesting costs.

There are several cases where the deadwood enhancement shouldbe avoided or carefully planned due to health and safety reasons. Theretention should be restricted near frequently used routes such asmarked hiking and cycling trails, where the retained or created dead-wood may present an increased risk to the safety of forest visitors.Another such example is where the tree retention forms an obstacle toharvesting or tending operations or transport routes. Tree retentionshould be carefully planned if the retained tree or tree group is locatedimmediately next to a fenced area in order to avoid or minimise damageto the fence due to a fall or a breakage of a fragment of the retainedgroup. In addition, retention of tree species susceptible to insect out-breaks shall be limited; e.g. refrain from retaining Norway spruce (Piceaabies L. Karst.) near stands dominated by this commercially importantconiferous species due to its susceptibility to bark beetle (Ips typo-graphus Linnaeus, 1758) outbreak. We have to be cautious especially inthe case of larger amounts of early stages of deadwood in single speciesplantations of Norway spruce. However, this effect is much less pro-nounced in forests with mixed species composition of structurally het-erogeneous stands.

The key measure allowing single trees or groups of trees to reach theend of their life span does not have to be strictly applied under allcircumstances. In order to minimise the financial loss, priority shouldbe given to retention of trees whose preservation produces smallereconomic loss. If an ideal deadwood type - known to greatly enhance

a) The total deadwood volume in the forest stands (m3/ha-1) and b) the mean volume of deadwood (m3/ha-1) in

individual decays classes (detail on decay stages in Appendix 1).

2.7

9.112.1

0.0

5.0

10.0

15.0

20.0

25.0

Low stumps Larger DW Small DW

Dea

dwoo

d vo

lum

e (m

3 /ha-

1 )

Deadwood type

a)

3.5

15.3

4.01.0

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5.0

10.0

15.0

20.0

25.0

30.0

0 1 2 3Mea

n de

adw

ood

volu

me

(m3 /h

a-1 )

Decay class

b)

Fig. 2. Case study of the potential of deadwood management in traditional commercially managed Central European forest property – the University forest Kostelecnad Černými lesy, in central Bohemia, Czech Republic. (a) The total deadwood volume in the forest stands (m3/ha−1) and (b) the mean volume of deadwood (m3/ha−1) in individual decays classes (detail on decay stages in Appendix A).


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biodiversity - is absent from the forests and its creation is difficult,generating different types of deadwood that supports other species withdifferent demands is also favourable. Deadwood retention with aminimum financial loss may also be applied in areas where the accessby transport vehicles is difficult or not possible due to extreme slopes,rugged terrain or where the area of interest is enclosed by properties ofother owners. Another case when retention may cause little financialloss is when the trees subjected to retention are of poor quality (e.g.trees showing irregular growth or poor form). Trees that already bearcertain stage of decay or harbour microhabitats may also be subjectedto retention at little financial loss. Trees found on the fringes of farm-land not utilised for farming that are adjacent to forests shall also beconsidered for long-term retention in the light of deadwood formation.

Kostelec nad Černými lesy (6900 ha) is a sustainably managed forestthat is used here to exemplify deadwood volumes and types in a typicalcommercially-managed Central European forest property focusing onthe production of construction timber, fuel wood, chips, bark, andbiomass. Norway spruce dominates half of the forest area with addi-tional 18 and 11% accounting for Scots pine and European beech, re-spectively, with the reminder comprising of other broadleaved andconiferous species. Inventory data from 2011 showed that deadwoodvolume varied across the forest with the mean deadwood volumereaching 23.9m3/ha−1 (a) with the younger decay stage (b; no. 1,Appendix A) being the most commonly represented throughout theforest. However, considering the total deadwood volume of 23.9 m3/ha−1 may be misleading since only the category of ‘larger deadwood’(snags and lying deadwood with the length> 1m and with the dia-meter at a smaller end>7 cm) forming 9.1 m3/ha−1 can be consideredas a suitable habitat for saproxylic species considered in this paper. Thereminding 14.8 m3/ha−1 account for small stumps (< 15 cm in dia-meter at the height of 20–30 cm) and small deadwood (deadwood<7 cm in diameter at a smaller end) that do not form an appropriatehabitat for the saproxylic species targeted by the deadwood manage-ment approaches described here.

Given sufficient time and dedicated forest management efforts, it ispossible to achieve internationally recommended deadwood volumesi.e. 35m3/ha−1 (e.g. Müller and Bütler, 2010) in this typical CentralEuropean commercial forest, if the timber assortments of poor qualityof larger sizes are retained. However, it is necessary to aim for greaterdiversity of deadwood types, especially larger deadwood segments inorder to satisfy the saproxylic beetles requiring larger fragments ofdeadwood as opposed to retaining smaller deadwood segments; i.e.stumps and stems or branches< 7 cm in diameter.

5. Discussion

Developing deadwood management recommendations requires agood comprehension of the relationships between deadwood char-acteristics and the ecology of species depending on it as part of their lifecycle (Brin et al., 2011). Certain deadwood management guidelineswere created on institutional levels in some countries or presented ingrey literature written in local languages. However, only a limitednumber of publications reported on the outcomes of deadwood man-agement in Central Europe. Although we did not present any novelinsights into the topic of deadwood in production forests, we were thefirst to concisely compile the key basic deadwood enhancement re-commendations that are based on scientific findings and practical ex-perience in the light of introducing the concept of feasible deadwoodmanagement in production forests of Central Europe. The presentedconcept of deadwood management demonstrates the balance betweenthe deadwood management benefits on biodiversity and the im-plementation of necessary deadwood management approaches notcompromising operational costs.

We provided a set of simple deadwood management approacheswith a minimum management input mainly focusing on the enhance-ment or creation of natural features in managed forests facilitating – not

only – saproxylic species. Although the disadvantages of deadwoodmanagement are apparent (e.g. conflict with timber production),deadwood management is a relatively low maintenance approach. Ifindividual deadwood enhancing methods present certain costs (i.e. ar-tificial creation of deadwood), we believe such cost are sufficientlybalanced by the benefits of individual deadwood management ap-proaches on biodiversity. Since most forests include areas that are notparticularly suitable for timber production (e.g. waterlogged soil, dif-ficult access, poor quality trees), such areas can be subjected to dead-wood retention without additional harvesting costs involved as they aregood candidates to maintain structural attributes. However, deadwoodmanagement shall be applied in the whole forest in order to achieve thepresence of deadwood across the entire forest site. It is encouraging toinclude some form of deadwood management practices in the business-as-usual forest management in public forests but also in forest proper-ties owned by communities and private owners since little or no cost isinvolved.

The willingness of forest managers to apply deadwood managementand impartially explain to private forest owners the pros and cons ofthis approach influences the forest owners to opt for deadwood man-agement with the aim to enhance biodiversity. Private owners may bestimulated to use some form of deadwood management since thedeadwood management approaches included here require minimumcosts but provide considerable enhancement of biodiversity. Therefore,such approaches are unlikely to hinder the management aims focusingon timber production, which, in contrary, are frequently seen as thereason not to opt for deadwood management. Nonetheless, virtual di-dactic tools such as marteloscopes can be utilised for exercising parti-cular forestry practices (Kraus et al., 2018). Such virtual training fa-cilities allow forestry professionals to carry out particular managementapproaches (e.g. tree retention for deadwood creation) in order to seethe outcomes without the necessity to retain or cut the selected trees.Such exercise helps build confidence in using alternative or unfamiliarpractices.

Forestry in Central Europe is a conservative discipline with deeplyrooted use of traditional silvicultural practices, some of which aim toreduce heterogeneity in forests, which is, however, the key factorsupporting biodiversity. In order to include what appears to be thenovel management concepts integrating multiple ecosystem servicesinto traditional forest management - such as deadwood enhancement -the mind set of many foresters requires a substantial renaissance. This ismainly due to the conflict between practising traditional silvicultureand accepting, and consequently incorporating, new approaches thatsometimes override the traditional and widely accepted silviculturalconcepts. In other words, retaining trees reaching target diameter forharvesting in order to create deadwood may be considered as one of thecompromises between biodiversity enhancement (in a form of dead-wood creation) and traditional forest management approaches theforesters ought to accept.

Deadwood management may as well be considered a novel ap-proach for some foresters. Recent research on the adoption of newforest management practices revealed that implementation of in-novative approaches may be challenging due to overriding reliance onfamiliar and widely-recognised techniques and due to the lack ofagreement amongst forestry professionals on the choice of trees forspecific purpose (e.g. Vítková et al., 2016; Pommerening et al., 2018).Since such findings are also applicable to choice of trees to be retainedas a part of deadwood management, a full appreciation of multipleforest functions would help forest managers to be more open-mindedand include some form of deadwood enhancement to better accom-modate for a wider range of forest services. Since certain certificationstandards require some level of deadwood retention for the purpose ofbiodiversity conservation, enforcing certification standards to certainextent positively stimulates the use of deadwood management.

It is also important to note that the growing societal demands andincreasing awareness of public regarding the quality of environment


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have already made many forest managers to apply approaches effec-tively accommodating for multiple ecosystem services. Employingdeadwood management is believed to help achieve this by means ofbalancing biodiversity enhancement and timber production. Whilebiodiversity is generally positively evaluated by public, the actualpresence of deadwood does not always get such a positive appraisal.This is mainly attributed to aesthetical values of deadwood as highamounts of deadwood in the forest are not often viewed as aestheticallypleasing but instead as messy, chaotic or abandoned. Such emotionalaspects should be also kept in mind by the practising forest managerswhen implementing deadwood management strategies in order tominimise any potential conflicts with the members of public. In addi-tion, in poorer regions where firewood is considered an important en-ergy resource, the application of deadwood management may be seenas a waste of valuable energy resource.

Certain aspects of deadwood, such as its position or decay stage,that are related to the occurrence of iconic or easily recognised species,have been subjected to more research in comparison to e.g. economicaspects of deadwood management where only a small amount of datahas been available. It is therefore necessary to carry out economic as-sessment in order to gain evidence on economic feasibility of individualdeadwood management approaches. For instance, cost-benefit analysesproviding information on the retention of single trees or groups of treesor on the artificial creation of deadwood would allow us to apply themost convenient forest management approach suited to the particularsite that balances interventions aiming at biodiversity enhancementwith those aiming at generating revenue from timber production. Thiswould facilitate for the ‘win-win’ situation where we manage for bio-diversity at minimum cost; i.e. reduce harvesting cost by retaining treegroups located in places with difficult access.

Health and safety in forests is another important point to be con-sidered as a part of deadwood management; especially, since forestworkers are often in a close contact with deadwood that may injurethem. Individual deadwood management approaches shall be thereforewell planned and carefully executed in order to minimise any healthand safety risks. Suitable methodology and practice policies are to beapplied or developed and an appropriate risk assessment shall be car-ried out prior to undertaking any work that ought to be carefully su-pervised and guided, if necessary. In addition, the risk of any injury tothe members of public or damage of their property shall be also mini-mised. However, the risk of injuries to forest visitors is generally lowdue to any dangerous deadwood segments being removed from within aclose proximity to public paths and roads in the forests due to a regularassessment of public areas and issuing cautions by means of informativesign posts.

6. Conclusion

Incorporation of deadwood management into conventional forestmanagement in production forests is a feasible forest management ap-proach that facilitates for biodiversity maintenance while simulta-neously allowing for timber production. However, individual deadwoodmanagement approaches need to be carefully selected in order to suitthe given forest conditions and management aims without acceptingunnecessary compromises. Although deadwood is an important featuresupporting forest biodiversity, only a limited number of forestry prac-titioners in Central Europe have actively engaged in approaches thateffectively enhance deadwood to recommended levels in forests wherethe major aim is timber production. Therefore, we highlighted the mainpractically feasible deadwood management approaches that can beused by forest managers in the forests of Central Europe to enhancedeadwood diversity and consequently overall forest biodiversity. Themajor factors relating to deadwood management discussed in thispaper, i.e. retention of trees with microhabitats, size of retained trees,position and arrangement, and decay stage, were highlighted as theyshall be considered in order to carry out successful deadwood

management.As a part of effective deadwood management practice, it is neces-

sary to retain as much of already existing deadwood (e.g. snags, fallenor lying deadwood, or uprooted and snapped trees) as possible.Retaining at least certain proportion of deadwood resulting from nat-ural disturbances that is usually subjected to salvage logging is alsofavourable for enhancing volume but most importantly also for dead-wood diversity. Most conventionally used silvicultural practices inCentral European forests do not actively engage in deadwood creation.Therefore, incorporating the retention of preferably sun-exposed singletrees or tree groups by means of allowing single trees or tree groups toreach the end of their life span eventually developing into snags thatdecay and disintegrate into coarse woody debris is essential. In addi-tion, artificial creation of deadwood where practices such as girdling,felling and pulling, inoculation with fungal pathogens or combinationof these techniques are employed are also valuable options for the en-hancement of deadwood diversity and volume.

Since tree size, its arrangement and position, but also the decay rateand tree species affect deadwood diversity and volume, consideringthese key factors greatly aids the decision-making on the choice ofparticular deadwood management approach. Moreover, bearing inmind these factors allows us to ensure that, given time, recommendeddeadwood volume is achieved and that its composition is diverse with arange of deadwood types, sizes, and decay stages. Large deadwoodsegments, preferably standing and sun-exposed, for instance, shall bepresent in the forests since they have longer residence time and supportgreater number of saproxylic species. Heterogeneous spatial distribu-tion of deadwood of various types is also important in order to achievevariability in spatial arrangement of deadwood. The focus on deadwoodformed by native species is advocated in the light of native biodiversityenhancement.

Deadwood management presents some disadvantages on opera-tional/commercial level such as the potential loss of financial revenuefrom timber harvesting, possible health and safety hazards or the ne-cessity to employ only costly specially trained staff to carry out theoperations. Disadvantages on biological level nonetheless also persist;i.e. unsuitable location or position of deadwood, the presence of un-suitable tree species with a fast decay rate, long time required untilsuitable decay stage is reached or deadwood presents a potential sourceof pests and diseases. However, careful planning of deadwood man-agement is required in order to eliminate the disadvantages and securecost-effective choice of particular approach. Therefore, given carefulchoice of deadwood management approach and sufficient time, in thecase any disadvantages remain, they are believed to be compensated bythe positive effects individual deadwood management approaches haveon biodiversity.

Acknowledgement

Radek Bače was supported by project CIGA No. 20164310 andproject LTC17055 financed by the Czech Ministry of Education, Youthand Sport project. Miroslav Svoboda was funded by project EVA4.0 No.CZ.02.1.01/0.0/0.0/16_019/0000803 financed by the Czech Ministryof Education, Youth and Sport. We also wish to thank to Péter Ódor,Yoan Paillet and an anonymous reviewer for their valuable commentsthat improved this paper.

Appendix A

Decay classes used to demonstrate the volumes of deadwood foundin the forest of Kostelec nad Černými lesy. The decay classes are basedon the methodology of Marušák et al. (2009) that is used for deadwoodevaluation in the Czech Republic and is comparable to other decayclasses grading system such those used in Siitonen et al., 2000,Heilmann-Clausen (2001), Nordén and Paltto (2001), Ódor and vanHees (2004), and Müller-Using and Bartsch (2009), etc.


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0: The tree recently died; its stem either collapsed or it is stillstanding; the wood is fresh, hard, with an absence of rot; the entirestem is covered by bark; the diameter of the smallest twigs is< 1cm.1: The wood remains hard with fungi already appearing; the stem iseither in a contact with the ground or it slants without groundcontact as it is supported by its branches; majority of the stem iscovered by bark in most cases; the diameter of the smallest twigsis> 1 cm.2: The wood is soft in places with pieces of decaying wood disin-tegrating; only thick branches are present; the stem in one piece andin a contact with the ground incompletely copying the ground sur-face; large clusters of epiphytic vegetation cover the stem; bark isabsent in most cases.3: The wood is very soft and disintegrating; the stem lies in a closecontact with the ground intimately copying the ground surface; thesurface of the stem is covered with deep furrows with the presenceof epiphytes and fungi; bark is absent in most cases.

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